Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ doi:10.5194/fr-19-131-2016 © Author(s) 2016. CC Attribution 3.0 License. First occurrence of Panthera atrox (Felidae, Pantherinae) in the Mexican state of Hidalgo and a review of the record of felids from the Pleistocene of Mexico Victor Manuel Bravo-Cuevas1, Jaime Priego-Vargas2, Miguel Ángel Cabral-Perdomo1, and Marco Antonio Pineda Maldonado3 1Museo de Paleontología, Área Académica de Biología, Universidad Autónoma del Estado de Hidalgo, Ciudad del Conocimiento, Carretera Pachuca-Tulancingo km 4.5, CP 42184, Pachuca, Hidalgo, Mexico 2Área Académica de Biología, Doctorado en Biodiversidad y Conservación, Universidad Autónoma del Estado de Hidalgo, Ciudad del Conocimiento, Carretera Pachuca-Tulancingo km 4.5, CP 42184, Pachuca, Hidalgo, Mexico 3Área Académica de Biología, Maestría en Biodiversidad y Conservación, Universidad Autónoma del Estado de Hidalgo, Ciudad del Conocimiento, Carretera Pachuca-Tulancingo km 4.5, CP 42184, Pachuca, Hidalgo, Mexico Correspondence to: Victor Manuel Bravo-Cuevas ([email protected]) Received: 26 May 2016 – Revised: 18 June 2016 – Accepted: 1 July 2016 – Published: 20 July 2016 Abstract. Panthera atrox was a common large-sized cat in the mesopredator level, such as the dire wolf (Canis dirus ), North America during the late Pleistocene. An isolated lower which has been also reported in the area. The high diversity canine and a fifth metacarpal bone referable to this species of large herbivores recorded at southeastern Hidalgo, which were recovered from fluvial Quaternary deposits that outcrop in turn could represent potential prey of P. atrox, suggests in southeastern Hidalgo, central Mexico. Associated fossil that some areas that now are part of central Mexico were suit- material belonging to Bison indicates a Rancholabrean North able hunting sites for this large-sized cat. American Land Mammal Age; the age assignment is corrob- A review of the Pleistocene record of Felidae from Mex- orated by the presence of P. atrox. A comparative study with ico revealed that it encompasses 87.5 and 73.3 % of generic selected specimens of Panthera and Smilodon indicates that and specific diversity known for North America, respec- the Hidalgoan sample shares the following diagnostic fea- tively, including seven genera (Felis, Miracinonyx, Panthera, tures with P. atrox: a large, robust, and non-strongly curved Puma, Lynx, Leopardus, and Smilodon) and 11 species (Fe- lower canine; a large and relatively slender fifth metacarpal lis rexroadensis, Miracinonyx inexpectatus, Panthera atrox, with a well-developed projection on the palmar side at the Panthera onca, Puma concolor, Puma yagouaroundi, Lynx proximal end, narrow articulating surface for the unciform; rufus, Leopardus pardalis, Leopardus wiedii, Smilodon fa- a narrow notch on the articulating surface for the fourth talis, and Smilodon gracilis). The majority of these taxa have metacarpal; and a diaphysis that at the middle is oval in cross been reported from numerous late Pleistocene localities; in section. The record supplements the evidence of P. atrox in particular, Panthera atrox was relatively common and widely central Mexico and represents the first reported occurrence distributed across the Mexican territory. of this cat species in the state of Hidalgo. By the same token, the known geographic distribution of P. atrox in the Mexi- can territory suggests that it was relatively common in tem- perate areas of central Mexico between 19 and 24◦ N at an 1 Introduction altitudinal range from 1500 to 2250 m a.s.l. The large size (mean body mass of 300 kg) and hypercarnivorous adapta- The subfamily Pantherinae is a group of large-sized cats that tions of the American lion suggest it was the top predator originated in Asia (probably in some part of what is now of the mammalian community recorded at southeastern Hi- China) during the end of the Miocene and at the beginning of dalgo, displacing other members of the carnivore guild at the Pliocene (10.8–3.8 Ma) (Johnson et al., 2006; Davis et al., Published by Copernicus Publications on behalf of the Museum für Naturkunde Berlin. 132 V. M. Bravo-Cuevas: First occurrence of Panthera atrox 2010; Tseng et al., 2016). This group of felids includes six 2 Study area extant species in the genera Panthera (P. leo (African lion), P. tigris (tiger), P. onca (jaguar), and P. pardus (leopard)), The material considered here has been recovered from a lo- ◦ 0 00 Uncia (U. uncia (snow leopard)), and Neofelis (N. nebulosa cality formally known as El Barrio (HGO-47: 20 07 41 N, ◦ 0 00 (clouded leopard)) (Davis et al., 2010; King and Wallace, 98 56 02.7 W) in the southeastern region of Hidalgo, cen- 2014); all of these species are considered to be vulnerable, tral Mexico (Fig. 1a). In this region Tertiary to Quaternary endangered, and/or threatened (IUCN, 2015). Extinct pan- volcanic and volcano sedimentary rock units crop out (IN- therines include the Pleistocene species P. gombaszoegensis EGI, 1992). The fossil-bearing strata consist of heteroge- (eastern Africa), P. fossilis (Eurasia), P. youngi (northeastern neous clastic material, including clay, silt, sand, and scarce China), P. spelaea (Eurasia and Alaska), and P. atrox (North conglomeratic lenses, setting in a fluvial depositional envi- America) (Kurtén and Anderson, 1980; Lange, 2002; Kurtén, ronment (Fig. 1b). 2009; Tseng et al., 2016). The oldest known pantherines are The associated mammalian fauna includes dire wolves those referable to P. blytheae from the late Miocene–early (Canis dirus), giant armadillos (Glyptotherium floridanum), Pliocene of the Himalayan region (Tseng et al., 2016) and P. giant ground sloths (Paramylodon cf. harlani), llamas (Hemi- palaeosinensis from the Plio-Pleistocene of China (Mazák, auchenia gracilis and Camelops sp.), horses (Equus conver- 2010). sidens), pronghorns (Stockoceros conklingi and Capromeryx The late Pleistocene species P. atrox was a common large- minor), deer (Odocoileus cf. virginianus), bison (Bison sp.), sized cat in the North American subcontinent, which has gomphotheres (Cuvieronius sp.), and mammoths (Mam- been traditionally referred to as the American lion (Kurtén muthus sp.) (Bravo-Cuevas, 2001, 2002; Bravo-Cuevas et al., and Anderson, 1980). Ancestors of the American lion appar- 2009, 2011, 2012, 2013). There is evidence of material be- ently crossed from Asia to Alaska sometime during the Illi- longing to the genus Bison, including a horn core fragment noian glacial stage (191 000 to 130 000 years ago) (Lange, and an isolated tooth; the presence of this taxon is indica- 2002) at the beginning of the Rancholabrean North Ameri- tive of a Rancholabrean NALMA (Bell et al., 2004). The can Land Mammal Age (NALMA). Early representatives of biochronological age of the fossil assemblage is supported by P. atrox appeared during the Sangamonian interglacial stage, the presence of material belonging to Panthera atrox (present reaching areas of southern temperate North America in a rel- study). atively short time (ca. 125 000 years ago) (Kurtén and An- derson, 1980; Lange, 2002). In the Wisconsinan (75 000– 3 Materials and methods 10 000 years ago) at the end of the Rancholabrean NALMA, P. atrox spread across the United States and Mexico (Kurtén The sample consists of an isolated canine and a metacarpal and Anderson, 1980), and its southernmost occurrence has bone. The material is housed at the Sección de Macrover- been recorded in tropical areas of the Mexican territory tebrados, Museo de Paleontología, Universidad Autónoma (Montellano-Ballesteros and Carbot-Chanona, 2009; Carbot- del Estado de Hidalgo, México, with the catalogue numbers Chanona and Gómez-Pérez, 2014). UAHMP-4221 and UAHMP-4222, respectively. The phylogenetic status of Panthera atrox is controver- The anteroposterior and transverse diameters of the iso- sial, given that some authors have considered it to have been lated tooth were measured. The measurements of the closely related to jaguars (Simpson, 1941; Christiansen and metacarpal are those of Merriam and Stock (1932: table 62, Harris, 2009) and others that it was a subspecies of the ex- p. 134). Measurements in millimeters were taken using a dig- tant lion (Kurtén and Anderson, 1980). Recent studies on the ital caliper with a measuring range of 0–150 mm, a resolution phylogenetic relationship of P. atrox with other members of of 0.01 mm, and an accuracy of 0.003 mm. the genus Panthera indicate that it should be considered as a In order to assess the taxonomic identity of the dental and separate species closely related to lions (P. leo) and tigers (P. postcranial remains from Hidalgo, we performed a compara- tigris) (Barnett et al., 2009; Christiansen and Harris, 2009; tive study with selected specimens of Panthera atrox housed King and Wallace, 2014). at the George C. Page Museum, La Brea Discoveries, Mu- Information regarding the fossil material, description, and seum of Natural History of Los Angeles County (LACM), formal taxonomic characterization of Mexican specimens California, USA. Additional comparisons were done with referable to P. atrox is unknown in many instances. Hence, specimens belonging to the extant species P. leo (OCMP- the purpose of this study is to formally describe the dental 080) and P. tigris (OCMP-077) housed at the Osteologi- and postcranial material belonging to Panthera atrox from cal Collection of the Museo de Paleontología, Universidad the late Pleistocene of southeastern Hidalgo, central Mexico, Autónoma del Estado de Hidalgo. Furthermore, the Hidal- as well as to discuss some ecological and geographic aspects goan material was compared with specimens described and of the record. Furthermore, a review of the record of felids figured in Merriam and Stock (1932), Whitmore and Fos- from the Pleistocene of Mexico is provided. ter (1967), and Montellano-Ballesteros and Carbot-Chanona (2009). Foss. Rec., 19, 131–141, 2016 www.foss-rec.net/19/131/2016/ V. M. Bravo-Cuevas: First occurrence of Panthera atrox 133 Figure 1. (a) Index map showing the study area in southeastern Hidalgo, central Mexico; the capital of the state (Pachuca) and the late Pleistocene locality El Barrio (HGO-47) are depicted.